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Long-term dietary change in Atlantic and Mediterranean Iberia with the introduction of agriculture: a stable isotope perspective

Long-term dietary change in Atlantic and Mediterranean Iberia with the introduction of... The Neolithic expansion in the Iberian Peninsula is marked by the introduction of livestock and domesticated crops which modified subsistence strategies in an unprecedented manner. Bulk collagen stable carbon and nitrogen isotope analysis has been essential to track these changes, which have largely been discussed in relation to particular geographic areas or single case studies. This paper reviews the available isotope literature to provide a regional, long-term synthesis of dietary changes associated with the expansion of the Neolithic and the establishment of farming economy in the Iberian Peninsula. Bulk collagen stable carbon and nitrogen isotope values of 763 human individuals and 283 faunal remains from the Mesolithic to the Late Neolithic period in Iberia (ca.8000–3000 cal BC) were collated and analysed using a Bayesian mixing model. The results show that Mesolithic diets were isotopically diverse in both the Atlantic and Mediterranean regions of the Iberian Peninsula, and that a significant decrease in variability happened with the Neolithisation, culminating with the establishment of farming economies and reliance on terrestrial resources in the Late Neolithic. . . . . Keywords Bulk collagen Stable carbon and nitrogen isotopes Palaeodietary reconstructions Neolithic Iberian Peninsula Introduction geographic contexts to investigate the magnitude and dietary implications of this process. First, the Peninsula was one of the The spread of agriculture in Europe during the Early to Middle most densely populated regions in Europe during the Mesolithic, Holocene constitutes one of the most transformative and contro- especially along the Atlantic coast (Arias 1999). Second, its geo- versial socio-cultural processes in Prehistory. Coastal zones and graphic diversity has demanded distinct adaptive strategies by immediate inland areas of the Iberian Peninsula offer attractive prehistoric populations, in particular by those living along the Atlantic and the Mediterranean, coastal areas marked by con- trasting biological productivity and resource predictability. Electronic supplementary material The online version of this article Archaeological evidence shows that early farmers from the (https://doi.org/10.1007/s12520-018-0752-1) contains supplementary material, which is available to authorized users. Near East migrated across the Mediterranean Sea following a coastal route from the Aegean to the Iberian Peninsula, bringing * Miriam Cubas with them pottery, ground stone tools and domesticated plants mcubas.morera@gmail.com and animals, which marked the beginning of the Neolithic in these regions (Guilaine 2017). Ancient human DNA has shown BioArCh-Department of Archaeology, University of York, a clear Anatolian ancestry of early European farmers (Mathieson Wentworth Way, Heslington, York YO10 5DD, UK et al. 2015; Omrak et al. 2016), as well as strong genetic differ- Sociedad de Ciencias Aranzadi, Zorroagagaina 11, entiation between hunter-gatherers and early farmers (Skoglund E-20014 Donostia, San Sebastian, Spain et al. 2014; Lazaridis et al. 2014), supporting the archaeological Human Evolution, Department of Organismal Biology, Uppsala model of a maritime colonisation of Iberia in the 6th millennium University, Norbyv, 18C, SE-752 36 Uppsala, Sweden cal BC (Zilhão 2001; Isern et al. 2014; Martins et al. 2015). UNIARQ, Faculdade de Letras da Universidade de Lisboa, Alameda The introduction of farming, however, was not uniform across da Universidade, University of Lisbon, 1600-214 Lisbon, Portugal Iberia. The first domesticates are known for the Mediterranean Department of Archaeology, Universiteit Leiden, Van Steenis region of the Peninsula, consisting of cereals and livestock, and Building, Einsteinweg 2, 2333 CC Leiden, Netherlands 3826 Archaeol Anthropol Sci (2019) 11:3825–3836 have been radiocarbon dated to the mid-6th millennium cal BC paper are twofold: (i) provide a regional and long-term synthesis (Saña 2013; Martins et al. 2015; Pérez-Jordá et al. 2017). of dietary changes associated with the introduction, expansion Domesticates were introduced to the south-western Atlantic and establishment of farming economy in this region from ca. coast in the second half of the 6th millennium cal BC (Zilhão 8000 to 3000 cal BC; and (ii) offer a comprehensive isotopic 2001; Davis and Simões 2015; López-Dóriga and Simões 2015), database for addressing the dietary implications of early food while in the north Atlantic coast of Iberia, such as the Cantabrian production in the Iberian Peninsula. region, the earliest domesticated plants and animals were adopted much later, in the 5th millennium cal BC (Cubas et al. 2016). Moreover, there has been a long, contentious debate about Materials and methods the impact of the expansion of farming upon local forager 13 15 diets, where the application of stable isotope analysis has been We analysed the bone collagen δ Cand δ N values current- particularly fruitful. Consumer diets can be determined by ly available for human individuals from archaeological sites 13 12 measuring the stable isotopes of carbon ( C/ C) and nitro- dated between ca. 8000 and 3000 cal BC in the Iberian 15 14 gen ( N/ N) in consumer tissues, such as bone collagen, and Peninsula (SI 1). Archaeological sites were grouped in two comparing these with isotopic baselines for potential food main regions (Atlantic, Mediterranean) and in generic sources (van Klinken et al. 1994; Tykot 2002; Fischer et al. chrono-cultural periods: Mesolithic (ca.8000–5500 cal BC), 13 15 2007). The δ Cand δ N values of human bone collagen Early Neolithic (ca. 5500–4500 cal BC), Middle Neolithic have been successfully used to estimate the relative contribu- (ca. 4500–3500 cal BC) and Late Neolithic–Chalcolithic tion of marine versus C -based terrestrial resources in coastal (ca.3500–3000 cal BC) (Fig. 1). Samples considered in this populations due to distinctive isotopic signatures in these eco- study are accompanied by their contextual and geographical systems, allowing generic discriminations (Jennings et al. information, cultural attribution and sample identification 1997; Richards and Hedges 1999). Isotopic analyses across (sample code and type, age and sex) (SI 1). Our analysis did Atlantic coastal areas of Europe have shown a marked dietary not distinguish between coastal and inland sites since in some difference between Mesolithic foragers, with substantial con- areas, such as the Atlantic coast of Iberia, the existence of sumption of marine foodstuffs, and Neolithic farmers, whose estuarine ecosystems allows marine resources to be exploited diets were essentially based on terrestrial resources (Lubell at considerable distances from the coast (e.g. Van Der Schriek et al. 1994; Richards and Hedges 1999;Richards etal. 2003; et al. 2007; Vis et al. 2008). The quality of the published Bonsall et al. 2009; Schulting 2011). Similar results have been material is extremely variable and data criteria are rarely pre- reported for the Iberian Peninsula, although considerable var- sented with sufficient detail to allow a critical evaluation of iability has been observed at the regional scale (Fontanals- analytical procedures (Roberts et al. 2018). Samples were Coll et al. 2014; Guiry et al. 2015; Peyroteo-Stjerna 2016; therefore screened according to established quality criteria Salazar-García et al. 2018). Indeed, since the early 2000s, for collagen preservation. Samples with C:N ratios outside there has been an extraordinary increase in palaeodietary stud- the 2.9–3.6 range (van Klinken 1999; DeNiro 1985) were ies based on stable isotope analysis of carbon and nitrogen in discounted as well as samples published without reporting human bone collagen from Mesolithic and Neolithic popula- these parameters. For this reason, some geographic areas, such tions in this region (e.g. Cunha and Umbelino 2001;Arias as the Cantabrian region (Arias 2006, 2012), are not well 2006; García-Guixé et al. 2006; Roksandic 2006;Umbelino represented in our database. 13 15 2006; Umbelino et al. 2007; Arias and Schulting 2010; Lillios The δ Cand δ N values of faunal remains from prehis- et al. 2010; McClure et al. 2010; Carvalho and Petchey 2013; toric and historic archaeological sites in the Atlantic and Fernández-López de Pablo et al. 2013; Jackes et al. 2014; Mediterranean regions were compiled to establish regional 13 15 Fontanals-Coll et al. 2014, 2015, 2017; Salazar-García et al. stable isotope baselines (SI 2). Fauna δ Cand δ Nvalues 2014; Gibajaetal. 2015; Gibaja et al. 2016; Guiry et al. 2015, were grouped according to their habitat: marine (fish and 2016;Umbelino et al. 2015; Alt et al. 2016; Jackes and Lubell mammals) and terrestrial (ungulates). Because of the paucity 2016;Peyroteo-Stjerna 2016;Remolins etal. 2016; Waterman of marine isotope values for the Iberian Peninsula, the δ C et al. 2016; Fernández-Crespo and Schulting 2017;Salazar- and δ N values of fish and marine mammals from Mesolithic García et al. 2017; Sarasketa-Gartzia et al. 2018). and Early Neolithic contexts from the central Mediterranean This paper reviews the isotope literature reporting δ Cand (Mannino et al. 2012, 2015), Bronze Age in the Balearic δ N values of human individuals from archaeological sites dat- Islands (García-Guixé et al. 2010) and Late Medieval data ed to the Mesolithic, Early Neolithic, Middle Neolithic and Late from the Atlantic (López-Costas and Müldner 2016)have Neolithic–Chalcolithic in the Iberian Peninsula. A Bayesian been also included. mixing model (FRUITS; Fernandes 2015) was used to estimate Statistical analyses were performed using R package v3.4 the average relative contribution of terrestrial and marine re- (R Development Core Team 2013). Different statistical tests sources to dietary calories for each time period. The aims of this were performed to test the normality of the distributions: Archaeol Anthropol Sci (2019) 11:3825–3836 3827 Fig. 1 Iberian Peninsula and the location of archaeological sites with stable isotope measurements of human bone considered in this study. a Mesolithic; b Early Neolithic; c Middle Neolithic; d Late Neolithic–Chalcolithic Shapiro–Wilk (S-W, for statistical populations with less than than two samples (Kruskal-Wallis, K-W). A p value of 0.05 50 cases) and Lilliefors corrected Kolmogórov–Smirnov (K- (H acceptance with p ≤ 0.05) was used as the significance S, for populations with more than 50 individuals). The results threshold. show that the isotope data were not normally distributed in all For each time period (Mesolithic, Early Neolithic, cases (SI 3 and 4). Non-parametric tests were then applied to Middle Neolithic and Late Neolithic-Chalcolithic) and re- compare two samples (Wilcoxon-Mann-Whitney, W) or more gion (Atlantic, Mediterranean), the average percentage of 3828 Archaeol Anthropol Sci (2019) 11:3825–3836 dietary carbon (equivalent to caloric contribution or dry and mammals, 17%) (García-Guixé et al. 2006; Salazar- weight contribution) derived from marine and terrestrial García 2009, 2012; Salazar-García et al. 2014, 2017; resources was estimated using a Bayesian Mixing Model Fontanals-Coll et al. 2015; Fernández-Crespo and Schulting in FRUITS 3.0 (Fernandes et al. 2014). While δ N 2017; Navarrete et al. 2017). values in bone collagen only reflect the sources of dietary Terrestrial faunal remains from the Mediterranean re- proteins and the position of the consumer in the food gion have significantly higher δ C(W =2006; p value = chain due to relatively predictable dietary isotope frac- 5.614e−05) and δ N(W = 1862; p value = 1.012e−05) tionations (+ 3–6‰;Jim et al. 2006;O’Connelletal. values compared to animals from the Atlantic region 2012), the δ C values may reflect the contribution of (Table 1). While no significant differences were observed different dietary macronutrients, such as proteins, carbo- for the δ C values of the marine animals between these hydrates and lipids, to individual diets (Jim et al. 2006; regions (W =94; p value = 0.1737), the fish and sea mam- Webb et al. 2017). A routed, concentration-dependent mals from the Atlantic region have δ N values signifi- weighted model was used taking into account two dietary cantly higher than those of specimens from the 13 15 proxies (δ C, δ N) and two food groups (marine, terres- Mediterranean (W = 244; p value = 0.002). This presum- trial). Isotopic baselines (Mediterranean and Atlantic) ably reflects fundamental differences in food webs be- were derived from published isotope data from animal tween the Mediterranean Sea compared to the Atlantic bones and are reported in Table 2. Although terrestrial Ocean (Stambler 2014). plants were undoubtedly an important portion of the diet The significant variability observed in the isotope of Mesolithic and Neolithic populations (e.g. Umbelino values of faunal remains from the Mediterranean and the 2006; López-Dóriga et al. 2016; Peña-Chocarro et al. Atlantic regions emphasises the importance of building 2018), the absence of isotopic values for both wild and local to regional baselines to estimate the contribution of 13 15 domestic plants in the region prevents their quantification. each food source. The average δ Cand δ Nvaluesof The model was developed using the assumptions and terrestrial and marine resources for each region were used uncertainties exactly as described by Fernandes (2015)as to establish the isotopic composition of macronutrients of this considers the dietary contributions of protein and en- each food group and the parameters required for model- ergy macronutrients. Rather than modelling individuals ling (Table 2). 13 15 separately based on their collagen δ Cand δ Nvalues, we aimed for broad comparisons by using the population Human bone collagen means for each region and each period, and by taking the standard deviations of the means as the uncertainties A total of 763 individuals were analysed from sites in the (Fernandes et al. 2015). Atlantic (n = 481) and Mediterranean regions (n = 282) (Fig. 1 and Table 3). Most individuals were dated to the Middle Neolithic (n = 315), followed by the Late Neolithic– Results Chalcolithic (n = 276), Mesolithic (n =150) and Early Neolithic (n = 22). Information about age and sex were not al- Fauna bone collagen ways available, preventing further analysis across populations. 13 15 The δ Cand δ N values show significant variability over 13 15 Faunal δ Cand δ N values are represented by 283 speci- time within and between regions (Fig. 2a, b). The largest range 13 15 mens from the Atlantic (n = 42) and Mediterranean (n =241) of δ Cand δ N values is observed among Mesolithic indi- regions (Table 1), with a disproportionate amount of terrestrial viduals of the Atlantic region. Although the paucity of isotope mammals (ungulates, 83%) compared to marine animals (fish values from the Early Neolithic prevents robust comparisons 13 15 Table 1 Faunal δ Cand δ N values grouped into two categories: terrestrial (ungulates), marine (fish and mammals) 13 15 δ C(‰) δ N(‰) N Mean SD Max Min Mean SD Max Min Atlantic Terrestrial − 20.8 1 − 18.9 − 22.1 4.3 1.6 9.6 2.1 35 Marine − 12 0.6 − 11.5 − 13 12.5 2.03 15.1 9.2 7 Mediterranean Terrestrial − 20 1.1 − 11.4 − 21.9 5.3 1.4 9 2.6 200 Marine − 11.6 1.6 − 8.9 − 15.2 9.2 2.1 12.1 3.6 41 Archaeol Anthropol Sci (2019) 11:3825–3836 3829 Table 2 Average and standard deviations of terrestrial and marine faunal collagen isotope values. The estimated protein and lipid isotope values and the macronutrient composition (%dry weight carbon content) were calculated according to Fernandes (2015) 13 15 13 δ C dietary protein (‰) δ N dietary protein (‰) δ C dietary lipids (‰) Protein (%C) Lipids (%C) Atlantic Terrestrial − 22.8 ± 1 6.2 ± 1 − 28.8 ± 1 30 ± 2.5 70 ± 2.5 Marine − 13 ± 1 14.5 ± 1 − 19 ± 1 65 ± 5 35 ± 5 Mediterranean Terrestrial fauna − 22 ± 1 7.3 ± 1 − 28 ± 1 30 ± 2.5 70 ± 2.5 Marine fauna − 12.6 ± 1 11.2 ± 1 − 18.6 ± 1 65 ± 5 35 ± 5 across regions for this time period (e.g. Mediterranean), there Atlantic region, Bayesian modelling estimates that terres- 13 15 is a decrease in δ Cand δ N variability in the Atlantic re- trial and marine resources provided average contributions gion. The narrow isotopic variability continued through the to dietary calories of 91.8 ± 6.2% and 8.2 ± 6.2%, respec- Middle to Late Neolithic–Chalcolithic, and these differences tively. However, marine resources may have contributed were statistically significant for both δ C (K-W = 172.81, as much as ca. 23% of dietary calories in some areas. df = 3, p value < 2.2e−16) and δ N values (K-W = 130.53, Indeed, considerable intake of marine resources has been df = 3, p value < 2.2e–16). In the Mediterranean region, reported for individuals from sites such as Cabeço do Pez, Mesolithic and Middle Neolithic populations had the most Cabeço da Amoreira, Cabeço da Arruda and Moita do 13 15 variable δ Cand δ N values, while considerably narrower Sebastião in Portugal (e.g. Lubell et al. 1994; Fontanals- isotopic distributions were recorded during the Late Coll et al. 2014; Guiry et al. 2015; Peyroteo-Stjerna Neolithic–Chalcolithic. These differences were also statisti- 2016). Thewiderangeof δ C values in the Atlantic cally significant for both δ C (K-W = 120.25, df = 3, p value region also suggests that distinct terrestrial ecological <2.2e–16) and δ N values (K-W = 7.9449, df = 3, p value = niches were exploited (e.g. Waterman 2012; Fontanals- 0.04716). Worth noting is the lower variability in δ Cand Coll et al. 2014). For the Mediterranean region, the δ N values of Mesolithic populations in the Mediterranean Bayesian model estimates average contributions of terres- region compared to the Atlantic. These differences could be a trial and marine resources of 95.5 ± 3.7% and 4.5 ± 3.7%, consequence of sample size, but most likely reflect differences respectively, to dietary calories. The maximum estimated in the isotope ecology and diet of populations between these contribution of marine resources to diet was ca. 14% regions, with considerable consumption of marine resources (Table 4). A higher consumption of marine resources in by some Atlantic foragers, as also estimated by Bayesian this region was found at the sites of El Collado, Santa calculations. Maira and some individuals from Cingle del Mas Nou (García-Guixé et al. 2006; Salazar-García et al. 2014). Mesolithic (ca. 8000–5500 cal BC) Early Neolithic (ca.5500–4500 cal BC) Stable isotope values of Mesolithic populations indicate diets based mainly on C -terrestrial foods in both regions Isotopic values from the Atlantic and Mediterranean re- of the peninsula (Fig. 2a, b; Fig. 3a; Table 4). For the gions are consistent with diets based on C -terrestrial 13 15 Table 3 Human δ Cand δ N values grouped by period 13 15 δ C(‰) δ N(‰) Period Mean SD Max Min Mean SD Max Min Atlantic Mesolithic (n =124) − 18 1.4 − 14.8 − 20.8 10.7 1.6 14 6.7 Early Neolithic (n =20) − 19.6 1 − 15.3 − 20.2 8.9 1.8 16.5 7.9 Middle Neolithic (n =174) − 19.4 0.8 − 14.8 − 21.3 9.1 1.2 13.4 6.7 Late Neolithic–Chalcolithic (n =163) − 19.9 0.5 − 18 − 21.3 8.6 0.8 11.7 6.9 Mediterranean Mesolithic (n =26) − 18.4 0.5 − 17.5 − 19.5 9.5 1.2 12.8 7.6 Early Neolithic (n =2) − 19 0.6 − 18.5 − 19.4 9.3 1.5 10.3 8.2 Middle Neolithic (n =141) − 19.5 0.7 − 16.8 − 21 9.2 1 13.1 6.1 Late Neolithic–Chalcolithic (n = 113) − 20.1 0.3 − 19 − 20.8 9.3 0.5 10.6 8.1 3830 Archaeol Anthropol Sci (2019) 11:3825–3836 Archaeol Anthropol Sci (2019) 11:3825–3836 3831 13 15 Fig. 2 δ C and δ N variation in human bone collagen and faunal are represented by individuals from Cerca do Zambujal and remains in the Atlantic and Mediterranean regions of the Iberian Lagar I (Guiry et al. 2016; Jackes and Lubell 2016)in the Peninsula from the Mesolithic to the Late Neolithic Atlantic region, and two individuals from Tossal de les Basses (Salazar-García et al. 2016) and one individual from Can resources, with an overall decline in marine intake Gambús (Fontanals-Coll et al. 2015) in the Mediterranean area (Fig. 2c, d; Fig. 3a, b). For the Atlantic region, Bayesian (SI 1). estimations indicate that terrestrial and marine resources provided average contributions of 95.7 ± 3.7% and 4.3 ± Late Neolithic–Chalcolithic (ca. 3500–3000 cal BC) 3.7% of dietary calories respectively (Table 4). The model estimates that the maximum amount of carbon from ma- 13 15 The δ Cand δ N values from the Atlantic and Mediterranean rine resources was ca.14% (Table 4). One individual has 13 15 regions are narrowly distributed and fall at the end-point of C - considerably higher δ Cand δ N values (Fig. 2c; terrestrial resources (Fig. 2g, h). The Bayesian model estimates Samouqueira 1 in Portugal; Lubell et al. 1994), but its average contributions of terrestrial resources to dietary calories of cultural attribution is controversial (Zilhão 2000). 98.1 ± 1.08% for the Atlantic and 98.4 ± 1.5% for the Similar results were obtained for Early Neolithic individ- Mediterranean. Marine resources contributed less than 6–7% in uals from the Mediterranean sites, where Bayesian model both regions (Table 4,Fig. 3a, b). estimates that terrestrial and marine resources accounted for averages of 96.1 ± 3.3% and 3.9 ± 3.3% of dietary cal- ories, respectively, while the maximum contribution of marine resources was ca. 13% (Tab. 4). Discussion Middle Neolithic (ca. 4500–3500 cal BC) The results show significant differences in human diets over time, from the Mesolithic to the Late Neolithic–Chalcolithic, Again, most of the samples have isotopic compositions consis- while they also varied geographically across the Atlantic and tent with C -terrestrial resources. This is supported by Bayesian Mediterranean regions of the Iberian Peninsula. The distribution 13 15 estimations of the terrestrial mammal contribution to dietary cal- of δ Cand δ N values and Bayesian estimations for Mesolithic ories with average values of 96.8 ± 2.7% for the Atlantic and populations reveal diets largely dominated by C -terrestrial re- 97.4 ± 2.4% for the Mediterranean (Table 4). Although the sources, here represented by ungulates. Terrestrial resources in- Middle Neolithic shows a clear shift toward a terrestrial-based cluding both animals and plants were the most significant com- diet, marine resources were consumed at a detectable level in ponent of the diet, in agreement with other lines of evidence several areas of the Peninsula (Fig. 2e, f). The average contribu- (Umbelino et al. 2007;Marín 2013; López-Dóriga et al. 2016). tion of marine resources to dietary calories was 3.2 ± 2.7% and The noticeable isotopic variability also indicates highly diversi- 2.6 ± 2.4% in the Atlantic and Mediterranean regions respective- fied diets, with groups exploiting a variety of ecological patches ly, but intakes of marine products up to 9–10% of the diet may and environments (Arias 2006; Fontanals-Coll et al. 2014; have occurred in both regions (Table 4,Fig. 3a, b). These values Salazar-García et al. 2014; Peyroteo-Stjerna 2016). Marine Fig. 3 Estimated marine calorie intake for the Atlantic (a)and credible interval. The horizontal continuous line represents the Mediterranean (b) populations in each chronological period. Boxes estimated mean and the horizontal discontinuous line represents the represent a 68% credible interval while the whiskers represent a 95% estimated median 3832 Archaeol Anthropol Sci (2019) 11:3825–3836 Table 4 Carbon contribution (%) of terrestrial and marine resources to the diet for each population Terrestrial vs. marine total carbon contribution to diet (%) Mean SD 2.5pc Median 97.5pc 16pc 84pc Atlantic Terrestrial Mesolithic (n = 124) 91.8 6.2 76.8 93.1 99.7 85.6 97.9 Early Neolithic (n = 20) 95.7 3.7 86.2 96.7 99.9 92.4 99.1 Middle Neolithic (n = 174) 96.8 2.7 90 97.5 99.9 94.3 99.3 Late Neolithic–Chalcolithic (n = 163) 98.1 1.8 93.4 98.6 100 96.6 99.6 Marine Mesolithic (n = 124) 8.2 6.2 0.3 6.9 23.2 2.2 14.4 Early Neolithic (n = 20) 4.3 3.7 0.1 3.3 13.8 0.9 7.6 Middle Neolithic (n = 174) 3.2 2.7 0.1 2.5 10 0.7 5.7 Late Neolithic–Chalcolithic (n = 163) 1.9 1.8 0.1 1.4 6.6 0.4 3.4 Mediterranean Terrestrial Mesolithic (n = 26) 95.5 3.7 86.3 96.4 99.8 92.1 99 Early Neolithic (n = 2) 96.1 3.3 87.5 96.9 99.9 93.1 99.1 Middle Neolithic (n = 141) 97.4 2.4 91.3 98.1 99.9 95.3 99.5 Late Neolithic–Chalcolithic (n = 113) 98.4 1.5 94.4 98.8 100 97 99.7 Marine Mesolithic (n = 26) 4.5 3.7 0.2 3.6 13.8 1 7.9 Early Neolithic (n = 2) 3.9 3.3 0.1 3.1 12.5 0.9 7 Middle Neolithic (n = 141) 2.6 2.4 0.1 1.9 8.7 0.5 4.7 Late Neolithic–Chalcolithic (n = 113) 1.6 1.5 0 1.2 5.6 0.3 3 resources contributed up to 23% of dietary calories, particularly increasingly anthropogenic landscape where productive along the Atlantic coast. Consumption of marine resources by cycles (crops and livestock) were fully integrated. While Mesolithic populations in the Atlantic region was possibly facil- this meta-analysis does not provide qualitative informa- itated by the higher marine productivity and the larger tidal zones tion on which resources were consumed, the interpreta- for shellfish gathering compared to the Mediterranean (Fa 2008). tions are generally supported by archaeozoological re- A decline in the consumption of marine foods took place with cords showing an increasing frequency of domesticated the introduction of farming in the Early Neolithic, which is par- animals from the Early to the Late Neolithic– ticularly noticeable in the Atlantic region, although it is important Chalcolithic in the Iberian Peninsula (e.g. Altuna and to highlight the low number of human remains dated in the Early Mariezkurrena 2009; Valente and Carvalho 2014). The Neolithic in the Iberian Peninsula in comparison to those dated to isotopic distribution of the Late Neolithic–Chalcolithic the Mesolithic and later periods (Fig. 1b). The paucity of data therefore probably reflects the establishment and intensi- from Mediterranean regions prevents a meaningful comparison fication of food production systems as indirectly evi- with the Atlantic, but we can expect a similar pattern (Salazar- denced by structural changes in settlement patterns, inten- García et al. 2018), even though archaeozoological records show sification of regional exchange networks and enhanced that marine resources were still exploited as food in some areas technological capacity (Bernabeu et al. 2013). (e.g. Benito 2015). Middle Neolithic individuals show the con- sumption of marine resources within economic contexts domi- natedbyC -based terrestrial diets. Higher human δ Nvalues Conclusions during the Middle Neolithic in both Atlantic and Mediterranean 13 15 Iberia have also been attributed to the consumption of freshwater Built on a large dataset of δ Cand δ N values of human and resources (Carvalho et al. 2016; Fontanals-Coll et al. 2017), but faunal remains in the available literature, this study offers a these might alternatively indicate protein intake from livestock long-term, regional synthesis of the transition from foraging to raised intensively on food leftovers (such as pigs) and/or ma- farming in the Iberian Peninsula. From the Mesolithic to the nured plants (Albarella et al. 2007;Bogaardetal. 2007, 2013; Late Neolithic–Chalcolithic, humans have largely relied on Navarrete et al. 2017). C -based terrestrial resources for food complemented, in some A definitive rupture with foraging subsistence strate- coastal areas, with marine organisms. Considerable isotopic gies appears to have occurred during the Late Neolithic– variability among Mesolithic individuals could be associ- Chalcolithic. The narrow isotopic distribution of individ- ated with versatile and diversified subsistence strategies, ual diets, in particular in the Mediterranean region, sug- adapted to the mosaic of environmental conditions in the gests an intensification of farming practices in an Iberian Peninsula. The onset of farming triggered the Archaeol Anthropol Sci (2019) 11:3825–3836 3833 Benito JLP (2015) El consumo de moluscos marinos durante el Neolítico replacement of heterogeneous diets based on a broad antiguo en la región central del Mediterráneo peninsular. spectrum of resources—including terrestrial animals from Archaeofauna 23:207–222 different ecological niches, marine resources and wild Bernabeu J, Moreno A, Barton M (2013) Complex systems, social net- plants—to fully terrestrial diets dominated by farming works, and the evolution of social complexity in the east of Spain from the Neolithic to pre-Roman times. In: Berrocal MC, García- products (crops and livestock) under increasing human Sanjuán L, Gilman A (eds) The prehistory of Iberia. Debating Early control of their production cycles. Social Stratification and the State. Routledge, New York, pp 53–73 Bogaard A, Heaton THE, Poulton P, Merbach I (2007) The impact of Acknowledgements This paper is a contribution to the Marie manuring on nitrogen isotope ratios in cereals: archaeological impli- Sklodowska Curie research project First ceramics of Atlantic Europe: cations for reconstruction of diet and crop management practices. J manufacture and function (CerAM -IF 653354-) funded by the Archaeol Sci 34:335–343. https://doi.org/10.1016/j.jas.2006.04.009 European Commission. The authors wish to thank Peter Smith and Bogaard A, Frase R, Heaton T, Wallace M, Vaiglova P, Charles M, Jones Krista McGrath for their review of the manuscript. In addition, we would G, Evershed R, Styring A, Andersen N, Arbogast RM, Bartosiewicz like to thank the two anonymous reviewers for their suggestions and L, Gardeisen A, Kanstrup M, Maier U, Marinova E, Ninov L, comments. Schäfer M, Stephan E (2013) Crop manuring and intensive land management by Europe’s first farmers. 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Long-term dietary change in Atlantic and Mediterranean Iberia with the introduction of agriculture: a stable isotope perspective

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References (114)

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Springer Journals
Copyright
Copyright © 2018 by The Author(s)
Subject
Earth Sciences; Earth Sciences, general; Archaeology; Chemistry/Food Science, general; Geography, general; Life Sciences, general; Anthropology
ISSN
1866-9557
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1866-9565
DOI
10.1007/s12520-018-0752-1
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Abstract

The Neolithic expansion in the Iberian Peninsula is marked by the introduction of livestock and domesticated crops which modified subsistence strategies in an unprecedented manner. Bulk collagen stable carbon and nitrogen isotope analysis has been essential to track these changes, which have largely been discussed in relation to particular geographic areas or single case studies. This paper reviews the available isotope literature to provide a regional, long-term synthesis of dietary changes associated with the expansion of the Neolithic and the establishment of farming economy in the Iberian Peninsula. Bulk collagen stable carbon and nitrogen isotope values of 763 human individuals and 283 faunal remains from the Mesolithic to the Late Neolithic period in Iberia (ca.8000–3000 cal BC) were collated and analysed using a Bayesian mixing model. The results show that Mesolithic diets were isotopically diverse in both the Atlantic and Mediterranean regions of the Iberian Peninsula, and that a significant decrease in variability happened with the Neolithisation, culminating with the establishment of farming economies and reliance on terrestrial resources in the Late Neolithic. . . . . Keywords Bulk collagen Stable carbon and nitrogen isotopes Palaeodietary reconstructions Neolithic Iberian Peninsula Introduction geographic contexts to investigate the magnitude and dietary implications of this process. First, the Peninsula was one of the The spread of agriculture in Europe during the Early to Middle most densely populated regions in Europe during the Mesolithic, Holocene constitutes one of the most transformative and contro- especially along the Atlantic coast (Arias 1999). Second, its geo- versial socio-cultural processes in Prehistory. Coastal zones and graphic diversity has demanded distinct adaptive strategies by immediate inland areas of the Iberian Peninsula offer attractive prehistoric populations, in particular by those living along the Atlantic and the Mediterranean, coastal areas marked by con- trasting biological productivity and resource predictability. Electronic supplementary material The online version of this article Archaeological evidence shows that early farmers from the (https://doi.org/10.1007/s12520-018-0752-1) contains supplementary material, which is available to authorized users. Near East migrated across the Mediterranean Sea following a coastal route from the Aegean to the Iberian Peninsula, bringing * Miriam Cubas with them pottery, ground stone tools and domesticated plants mcubas.morera@gmail.com and animals, which marked the beginning of the Neolithic in these regions (Guilaine 2017). Ancient human DNA has shown BioArCh-Department of Archaeology, University of York, a clear Anatolian ancestry of early European farmers (Mathieson Wentworth Way, Heslington, York YO10 5DD, UK et al. 2015; Omrak et al. 2016), as well as strong genetic differ- Sociedad de Ciencias Aranzadi, Zorroagagaina 11, entiation between hunter-gatherers and early farmers (Skoglund E-20014 Donostia, San Sebastian, Spain et al. 2014; Lazaridis et al. 2014), supporting the archaeological Human Evolution, Department of Organismal Biology, Uppsala model of a maritime colonisation of Iberia in the 6th millennium University, Norbyv, 18C, SE-752 36 Uppsala, Sweden cal BC (Zilhão 2001; Isern et al. 2014; Martins et al. 2015). UNIARQ, Faculdade de Letras da Universidade de Lisboa, Alameda The introduction of farming, however, was not uniform across da Universidade, University of Lisbon, 1600-214 Lisbon, Portugal Iberia. The first domesticates are known for the Mediterranean Department of Archaeology, Universiteit Leiden, Van Steenis region of the Peninsula, consisting of cereals and livestock, and Building, Einsteinweg 2, 2333 CC Leiden, Netherlands 3826 Archaeol Anthropol Sci (2019) 11:3825–3836 have been radiocarbon dated to the mid-6th millennium cal BC paper are twofold: (i) provide a regional and long-term synthesis (Saña 2013; Martins et al. 2015; Pérez-Jordá et al. 2017). of dietary changes associated with the introduction, expansion Domesticates were introduced to the south-western Atlantic and establishment of farming economy in this region from ca. coast in the second half of the 6th millennium cal BC (Zilhão 8000 to 3000 cal BC; and (ii) offer a comprehensive isotopic 2001; Davis and Simões 2015; López-Dóriga and Simões 2015), database for addressing the dietary implications of early food while in the north Atlantic coast of Iberia, such as the Cantabrian production in the Iberian Peninsula. region, the earliest domesticated plants and animals were adopted much later, in the 5th millennium cal BC (Cubas et al. 2016). Moreover, there has been a long, contentious debate about Materials and methods the impact of the expansion of farming upon local forager 13 15 diets, where the application of stable isotope analysis has been We analysed the bone collagen δ Cand δ N values current- particularly fruitful. Consumer diets can be determined by ly available for human individuals from archaeological sites 13 12 measuring the stable isotopes of carbon ( C/ C) and nitro- dated between ca. 8000 and 3000 cal BC in the Iberian 15 14 gen ( N/ N) in consumer tissues, such as bone collagen, and Peninsula (SI 1). Archaeological sites were grouped in two comparing these with isotopic baselines for potential food main regions (Atlantic, Mediterranean) and in generic sources (van Klinken et al. 1994; Tykot 2002; Fischer et al. chrono-cultural periods: Mesolithic (ca.8000–5500 cal BC), 13 15 2007). The δ Cand δ N values of human bone collagen Early Neolithic (ca. 5500–4500 cal BC), Middle Neolithic have been successfully used to estimate the relative contribu- (ca. 4500–3500 cal BC) and Late Neolithic–Chalcolithic tion of marine versus C -based terrestrial resources in coastal (ca.3500–3000 cal BC) (Fig. 1). Samples considered in this populations due to distinctive isotopic signatures in these eco- study are accompanied by their contextual and geographical systems, allowing generic discriminations (Jennings et al. information, cultural attribution and sample identification 1997; Richards and Hedges 1999). Isotopic analyses across (sample code and type, age and sex) (SI 1). Our analysis did Atlantic coastal areas of Europe have shown a marked dietary not distinguish between coastal and inland sites since in some difference between Mesolithic foragers, with substantial con- areas, such as the Atlantic coast of Iberia, the existence of sumption of marine foodstuffs, and Neolithic farmers, whose estuarine ecosystems allows marine resources to be exploited diets were essentially based on terrestrial resources (Lubell at considerable distances from the coast (e.g. Van Der Schriek et al. 1994; Richards and Hedges 1999;Richards etal. 2003; et al. 2007; Vis et al. 2008). The quality of the published Bonsall et al. 2009; Schulting 2011). Similar results have been material is extremely variable and data criteria are rarely pre- reported for the Iberian Peninsula, although considerable var- sented with sufficient detail to allow a critical evaluation of iability has been observed at the regional scale (Fontanals- analytical procedures (Roberts et al. 2018). Samples were Coll et al. 2014; Guiry et al. 2015; Peyroteo-Stjerna 2016; therefore screened according to established quality criteria Salazar-García et al. 2018). Indeed, since the early 2000s, for collagen preservation. Samples with C:N ratios outside there has been an extraordinary increase in palaeodietary stud- the 2.9–3.6 range (van Klinken 1999; DeNiro 1985) were ies based on stable isotope analysis of carbon and nitrogen in discounted as well as samples published without reporting human bone collagen from Mesolithic and Neolithic popula- these parameters. For this reason, some geographic areas, such tions in this region (e.g. Cunha and Umbelino 2001;Arias as the Cantabrian region (Arias 2006, 2012), are not well 2006; García-Guixé et al. 2006; Roksandic 2006;Umbelino represented in our database. 13 15 2006; Umbelino et al. 2007; Arias and Schulting 2010; Lillios The δ Cand δ N values of faunal remains from prehis- et al. 2010; McClure et al. 2010; Carvalho and Petchey 2013; toric and historic archaeological sites in the Atlantic and Fernández-López de Pablo et al. 2013; Jackes et al. 2014; Mediterranean regions were compiled to establish regional 13 15 Fontanals-Coll et al. 2014, 2015, 2017; Salazar-García et al. stable isotope baselines (SI 2). Fauna δ Cand δ Nvalues 2014; Gibajaetal. 2015; Gibaja et al. 2016; Guiry et al. 2015, were grouped according to their habitat: marine (fish and 2016;Umbelino et al. 2015; Alt et al. 2016; Jackes and Lubell mammals) and terrestrial (ungulates). Because of the paucity 2016;Peyroteo-Stjerna 2016;Remolins etal. 2016; Waterman of marine isotope values for the Iberian Peninsula, the δ C et al. 2016; Fernández-Crespo and Schulting 2017;Salazar- and δ N values of fish and marine mammals from Mesolithic García et al. 2017; Sarasketa-Gartzia et al. 2018). and Early Neolithic contexts from the central Mediterranean This paper reviews the isotope literature reporting δ Cand (Mannino et al. 2012, 2015), Bronze Age in the Balearic δ N values of human individuals from archaeological sites dat- Islands (García-Guixé et al. 2010) and Late Medieval data ed to the Mesolithic, Early Neolithic, Middle Neolithic and Late from the Atlantic (López-Costas and Müldner 2016)have Neolithic–Chalcolithic in the Iberian Peninsula. A Bayesian been also included. mixing model (FRUITS; Fernandes 2015) was used to estimate Statistical analyses were performed using R package v3.4 the average relative contribution of terrestrial and marine re- (R Development Core Team 2013). Different statistical tests sources to dietary calories for each time period. The aims of this were performed to test the normality of the distributions: Archaeol Anthropol Sci (2019) 11:3825–3836 3827 Fig. 1 Iberian Peninsula and the location of archaeological sites with stable isotope measurements of human bone considered in this study. a Mesolithic; b Early Neolithic; c Middle Neolithic; d Late Neolithic–Chalcolithic Shapiro–Wilk (S-W, for statistical populations with less than than two samples (Kruskal-Wallis, K-W). A p value of 0.05 50 cases) and Lilliefors corrected Kolmogórov–Smirnov (K- (H acceptance with p ≤ 0.05) was used as the significance S, for populations with more than 50 individuals). The results threshold. show that the isotope data were not normally distributed in all For each time period (Mesolithic, Early Neolithic, cases (SI 3 and 4). Non-parametric tests were then applied to Middle Neolithic and Late Neolithic-Chalcolithic) and re- compare two samples (Wilcoxon-Mann-Whitney, W) or more gion (Atlantic, Mediterranean), the average percentage of 3828 Archaeol Anthropol Sci (2019) 11:3825–3836 dietary carbon (equivalent to caloric contribution or dry and mammals, 17%) (García-Guixé et al. 2006; Salazar- weight contribution) derived from marine and terrestrial García 2009, 2012; Salazar-García et al. 2014, 2017; resources was estimated using a Bayesian Mixing Model Fontanals-Coll et al. 2015; Fernández-Crespo and Schulting in FRUITS 3.0 (Fernandes et al. 2014). While δ N 2017; Navarrete et al. 2017). values in bone collagen only reflect the sources of dietary Terrestrial faunal remains from the Mediterranean re- proteins and the position of the consumer in the food gion have significantly higher δ C(W =2006; p value = chain due to relatively predictable dietary isotope frac- 5.614e−05) and δ N(W = 1862; p value = 1.012e−05) tionations (+ 3–6‰;Jim et al. 2006;O’Connelletal. values compared to animals from the Atlantic region 2012), the δ C values may reflect the contribution of (Table 1). While no significant differences were observed different dietary macronutrients, such as proteins, carbo- for the δ C values of the marine animals between these hydrates and lipids, to individual diets (Jim et al. 2006; regions (W =94; p value = 0.1737), the fish and sea mam- Webb et al. 2017). A routed, concentration-dependent mals from the Atlantic region have δ N values signifi- weighted model was used taking into account two dietary cantly higher than those of specimens from the 13 15 proxies (δ C, δ N) and two food groups (marine, terres- Mediterranean (W = 244; p value = 0.002). This presum- trial). Isotopic baselines (Mediterranean and Atlantic) ably reflects fundamental differences in food webs be- were derived from published isotope data from animal tween the Mediterranean Sea compared to the Atlantic bones and are reported in Table 2. Although terrestrial Ocean (Stambler 2014). plants were undoubtedly an important portion of the diet The significant variability observed in the isotope of Mesolithic and Neolithic populations (e.g. Umbelino values of faunal remains from the Mediterranean and the 2006; López-Dóriga et al. 2016; Peña-Chocarro et al. Atlantic regions emphasises the importance of building 2018), the absence of isotopic values for both wild and local to regional baselines to estimate the contribution of 13 15 domestic plants in the region prevents their quantification. each food source. The average δ Cand δ Nvaluesof The model was developed using the assumptions and terrestrial and marine resources for each region were used uncertainties exactly as described by Fernandes (2015)as to establish the isotopic composition of macronutrients of this considers the dietary contributions of protein and en- each food group and the parameters required for model- ergy macronutrients. Rather than modelling individuals ling (Table 2). 13 15 separately based on their collagen δ Cand δ Nvalues, we aimed for broad comparisons by using the population Human bone collagen means for each region and each period, and by taking the standard deviations of the means as the uncertainties A total of 763 individuals were analysed from sites in the (Fernandes et al. 2015). Atlantic (n = 481) and Mediterranean regions (n = 282) (Fig. 1 and Table 3). Most individuals were dated to the Middle Neolithic (n = 315), followed by the Late Neolithic– Results Chalcolithic (n = 276), Mesolithic (n =150) and Early Neolithic (n = 22). Information about age and sex were not al- Fauna bone collagen ways available, preventing further analysis across populations. 13 15 The δ Cand δ N values show significant variability over 13 15 Faunal δ Cand δ N values are represented by 283 speci- time within and between regions (Fig. 2a, b). The largest range 13 15 mens from the Atlantic (n = 42) and Mediterranean (n =241) of δ Cand δ N values is observed among Mesolithic indi- regions (Table 1), with a disproportionate amount of terrestrial viduals of the Atlantic region. Although the paucity of isotope mammals (ungulates, 83%) compared to marine animals (fish values from the Early Neolithic prevents robust comparisons 13 15 Table 1 Faunal δ Cand δ N values grouped into two categories: terrestrial (ungulates), marine (fish and mammals) 13 15 δ C(‰) δ N(‰) N Mean SD Max Min Mean SD Max Min Atlantic Terrestrial − 20.8 1 − 18.9 − 22.1 4.3 1.6 9.6 2.1 35 Marine − 12 0.6 − 11.5 − 13 12.5 2.03 15.1 9.2 7 Mediterranean Terrestrial − 20 1.1 − 11.4 − 21.9 5.3 1.4 9 2.6 200 Marine − 11.6 1.6 − 8.9 − 15.2 9.2 2.1 12.1 3.6 41 Archaeol Anthropol Sci (2019) 11:3825–3836 3829 Table 2 Average and standard deviations of terrestrial and marine faunal collagen isotope values. The estimated protein and lipid isotope values and the macronutrient composition (%dry weight carbon content) were calculated according to Fernandes (2015) 13 15 13 δ C dietary protein (‰) δ N dietary protein (‰) δ C dietary lipids (‰) Protein (%C) Lipids (%C) Atlantic Terrestrial − 22.8 ± 1 6.2 ± 1 − 28.8 ± 1 30 ± 2.5 70 ± 2.5 Marine − 13 ± 1 14.5 ± 1 − 19 ± 1 65 ± 5 35 ± 5 Mediterranean Terrestrial fauna − 22 ± 1 7.3 ± 1 − 28 ± 1 30 ± 2.5 70 ± 2.5 Marine fauna − 12.6 ± 1 11.2 ± 1 − 18.6 ± 1 65 ± 5 35 ± 5 across regions for this time period (e.g. Mediterranean), there Atlantic region, Bayesian modelling estimates that terres- 13 15 is a decrease in δ Cand δ N variability in the Atlantic re- trial and marine resources provided average contributions gion. The narrow isotopic variability continued through the to dietary calories of 91.8 ± 6.2% and 8.2 ± 6.2%, respec- Middle to Late Neolithic–Chalcolithic, and these differences tively. However, marine resources may have contributed were statistically significant for both δ C (K-W = 172.81, as much as ca. 23% of dietary calories in some areas. df = 3, p value < 2.2e−16) and δ N values (K-W = 130.53, Indeed, considerable intake of marine resources has been df = 3, p value < 2.2e–16). In the Mediterranean region, reported for individuals from sites such as Cabeço do Pez, Mesolithic and Middle Neolithic populations had the most Cabeço da Amoreira, Cabeço da Arruda and Moita do 13 15 variable δ Cand δ N values, while considerably narrower Sebastião in Portugal (e.g. Lubell et al. 1994; Fontanals- isotopic distributions were recorded during the Late Coll et al. 2014; Guiry et al. 2015; Peyroteo-Stjerna Neolithic–Chalcolithic. These differences were also statisti- 2016). Thewiderangeof δ C values in the Atlantic cally significant for both δ C (K-W = 120.25, df = 3, p value region also suggests that distinct terrestrial ecological <2.2e–16) and δ N values (K-W = 7.9449, df = 3, p value = niches were exploited (e.g. Waterman 2012; Fontanals- 0.04716). Worth noting is the lower variability in δ Cand Coll et al. 2014). For the Mediterranean region, the δ N values of Mesolithic populations in the Mediterranean Bayesian model estimates average contributions of terres- region compared to the Atlantic. These differences could be a trial and marine resources of 95.5 ± 3.7% and 4.5 ± 3.7%, consequence of sample size, but most likely reflect differences respectively, to dietary calories. The maximum estimated in the isotope ecology and diet of populations between these contribution of marine resources to diet was ca. 14% regions, with considerable consumption of marine resources (Table 4). A higher consumption of marine resources in by some Atlantic foragers, as also estimated by Bayesian this region was found at the sites of El Collado, Santa calculations. Maira and some individuals from Cingle del Mas Nou (García-Guixé et al. 2006; Salazar-García et al. 2014). Mesolithic (ca. 8000–5500 cal BC) Early Neolithic (ca.5500–4500 cal BC) Stable isotope values of Mesolithic populations indicate diets based mainly on C -terrestrial foods in both regions Isotopic values from the Atlantic and Mediterranean re- of the peninsula (Fig. 2a, b; Fig. 3a; Table 4). For the gions are consistent with diets based on C -terrestrial 13 15 Table 3 Human δ Cand δ N values grouped by period 13 15 δ C(‰) δ N(‰) Period Mean SD Max Min Mean SD Max Min Atlantic Mesolithic (n =124) − 18 1.4 − 14.8 − 20.8 10.7 1.6 14 6.7 Early Neolithic (n =20) − 19.6 1 − 15.3 − 20.2 8.9 1.8 16.5 7.9 Middle Neolithic (n =174) − 19.4 0.8 − 14.8 − 21.3 9.1 1.2 13.4 6.7 Late Neolithic–Chalcolithic (n =163) − 19.9 0.5 − 18 − 21.3 8.6 0.8 11.7 6.9 Mediterranean Mesolithic (n =26) − 18.4 0.5 − 17.5 − 19.5 9.5 1.2 12.8 7.6 Early Neolithic (n =2) − 19 0.6 − 18.5 − 19.4 9.3 1.5 10.3 8.2 Middle Neolithic (n =141) − 19.5 0.7 − 16.8 − 21 9.2 1 13.1 6.1 Late Neolithic–Chalcolithic (n = 113) − 20.1 0.3 − 19 − 20.8 9.3 0.5 10.6 8.1 3830 Archaeol Anthropol Sci (2019) 11:3825–3836 Archaeol Anthropol Sci (2019) 11:3825–3836 3831 13 15 Fig. 2 δ C and δ N variation in human bone collagen and faunal are represented by individuals from Cerca do Zambujal and remains in the Atlantic and Mediterranean regions of the Iberian Lagar I (Guiry et al. 2016; Jackes and Lubell 2016)in the Peninsula from the Mesolithic to the Late Neolithic Atlantic region, and two individuals from Tossal de les Basses (Salazar-García et al. 2016) and one individual from Can resources, with an overall decline in marine intake Gambús (Fontanals-Coll et al. 2015) in the Mediterranean area (Fig. 2c, d; Fig. 3a, b). For the Atlantic region, Bayesian (SI 1). estimations indicate that terrestrial and marine resources provided average contributions of 95.7 ± 3.7% and 4.3 ± Late Neolithic–Chalcolithic (ca. 3500–3000 cal BC) 3.7% of dietary calories respectively (Table 4). The model estimates that the maximum amount of carbon from ma- 13 15 The δ Cand δ N values from the Atlantic and Mediterranean rine resources was ca.14% (Table 4). One individual has 13 15 regions are narrowly distributed and fall at the end-point of C - considerably higher δ Cand δ N values (Fig. 2c; terrestrial resources (Fig. 2g, h). The Bayesian model estimates Samouqueira 1 in Portugal; Lubell et al. 1994), but its average contributions of terrestrial resources to dietary calories of cultural attribution is controversial (Zilhão 2000). 98.1 ± 1.08% for the Atlantic and 98.4 ± 1.5% for the Similar results were obtained for Early Neolithic individ- Mediterranean. Marine resources contributed less than 6–7% in uals from the Mediterranean sites, where Bayesian model both regions (Table 4,Fig. 3a, b). estimates that terrestrial and marine resources accounted for averages of 96.1 ± 3.3% and 3.9 ± 3.3% of dietary cal- ories, respectively, while the maximum contribution of marine resources was ca. 13% (Tab. 4). Discussion Middle Neolithic (ca. 4500–3500 cal BC) The results show significant differences in human diets over time, from the Mesolithic to the Late Neolithic–Chalcolithic, Again, most of the samples have isotopic compositions consis- while they also varied geographically across the Atlantic and tent with C -terrestrial resources. This is supported by Bayesian Mediterranean regions of the Iberian Peninsula. The distribution 13 15 estimations of the terrestrial mammal contribution to dietary cal- of δ Cand δ N values and Bayesian estimations for Mesolithic ories with average values of 96.8 ± 2.7% for the Atlantic and populations reveal diets largely dominated by C -terrestrial re- 97.4 ± 2.4% for the Mediterranean (Table 4). Although the sources, here represented by ungulates. Terrestrial resources in- Middle Neolithic shows a clear shift toward a terrestrial-based cluding both animals and plants were the most significant com- diet, marine resources were consumed at a detectable level in ponent of the diet, in agreement with other lines of evidence several areas of the Peninsula (Fig. 2e, f). The average contribu- (Umbelino et al. 2007;Marín 2013; López-Dóriga et al. 2016). tion of marine resources to dietary calories was 3.2 ± 2.7% and The noticeable isotopic variability also indicates highly diversi- 2.6 ± 2.4% in the Atlantic and Mediterranean regions respective- fied diets, with groups exploiting a variety of ecological patches ly, but intakes of marine products up to 9–10% of the diet may and environments (Arias 2006; Fontanals-Coll et al. 2014; have occurred in both regions (Table 4,Fig. 3a, b). These values Salazar-García et al. 2014; Peyroteo-Stjerna 2016). Marine Fig. 3 Estimated marine calorie intake for the Atlantic (a)and credible interval. The horizontal continuous line represents the Mediterranean (b) populations in each chronological period. Boxes estimated mean and the horizontal discontinuous line represents the represent a 68% credible interval while the whiskers represent a 95% estimated median 3832 Archaeol Anthropol Sci (2019) 11:3825–3836 Table 4 Carbon contribution (%) of terrestrial and marine resources to the diet for each population Terrestrial vs. marine total carbon contribution to diet (%) Mean SD 2.5pc Median 97.5pc 16pc 84pc Atlantic Terrestrial Mesolithic (n = 124) 91.8 6.2 76.8 93.1 99.7 85.6 97.9 Early Neolithic (n = 20) 95.7 3.7 86.2 96.7 99.9 92.4 99.1 Middle Neolithic (n = 174) 96.8 2.7 90 97.5 99.9 94.3 99.3 Late Neolithic–Chalcolithic (n = 163) 98.1 1.8 93.4 98.6 100 96.6 99.6 Marine Mesolithic (n = 124) 8.2 6.2 0.3 6.9 23.2 2.2 14.4 Early Neolithic (n = 20) 4.3 3.7 0.1 3.3 13.8 0.9 7.6 Middle Neolithic (n = 174) 3.2 2.7 0.1 2.5 10 0.7 5.7 Late Neolithic–Chalcolithic (n = 163) 1.9 1.8 0.1 1.4 6.6 0.4 3.4 Mediterranean Terrestrial Mesolithic (n = 26) 95.5 3.7 86.3 96.4 99.8 92.1 99 Early Neolithic (n = 2) 96.1 3.3 87.5 96.9 99.9 93.1 99.1 Middle Neolithic (n = 141) 97.4 2.4 91.3 98.1 99.9 95.3 99.5 Late Neolithic–Chalcolithic (n = 113) 98.4 1.5 94.4 98.8 100 97 99.7 Marine Mesolithic (n = 26) 4.5 3.7 0.2 3.6 13.8 1 7.9 Early Neolithic (n = 2) 3.9 3.3 0.1 3.1 12.5 0.9 7 Middle Neolithic (n = 141) 2.6 2.4 0.1 1.9 8.7 0.5 4.7 Late Neolithic–Chalcolithic (n = 113) 1.6 1.5 0 1.2 5.6 0.3 3 resources contributed up to 23% of dietary calories, particularly increasingly anthropogenic landscape where productive along the Atlantic coast. Consumption of marine resources by cycles (crops and livestock) were fully integrated. While Mesolithic populations in the Atlantic region was possibly facil- this meta-analysis does not provide qualitative informa- itated by the higher marine productivity and the larger tidal zones tion on which resources were consumed, the interpreta- for shellfish gathering compared to the Mediterranean (Fa 2008). tions are generally supported by archaeozoological re- A decline in the consumption of marine foods took place with cords showing an increasing frequency of domesticated the introduction of farming in the Early Neolithic, which is par- animals from the Early to the Late Neolithic– ticularly noticeable in the Atlantic region, although it is important Chalcolithic in the Iberian Peninsula (e.g. Altuna and to highlight the low number of human remains dated in the Early Mariezkurrena 2009; Valente and Carvalho 2014). The Neolithic in the Iberian Peninsula in comparison to those dated to isotopic distribution of the Late Neolithic–Chalcolithic the Mesolithic and later periods (Fig. 1b). The paucity of data therefore probably reflects the establishment and intensi- from Mediterranean regions prevents a meaningful comparison fication of food production systems as indirectly evi- with the Atlantic, but we can expect a similar pattern (Salazar- denced by structural changes in settlement patterns, inten- García et al. 2018), even though archaeozoological records show sification of regional exchange networks and enhanced that marine resources were still exploited as food in some areas technological capacity (Bernabeu et al. 2013). (e.g. Benito 2015). Middle Neolithic individuals show the con- sumption of marine resources within economic contexts domi- natedbyC -based terrestrial diets. Higher human δ Nvalues Conclusions during the Middle Neolithic in both Atlantic and Mediterranean 13 15 Iberia have also been attributed to the consumption of freshwater Built on a large dataset of δ Cand δ N values of human and resources (Carvalho et al. 2016; Fontanals-Coll et al. 2017), but faunal remains in the available literature, this study offers a these might alternatively indicate protein intake from livestock long-term, regional synthesis of the transition from foraging to raised intensively on food leftovers (such as pigs) and/or ma- farming in the Iberian Peninsula. From the Mesolithic to the nured plants (Albarella et al. 2007;Bogaardetal. 2007, 2013; Late Neolithic–Chalcolithic, humans have largely relied on Navarrete et al. 2017). C -based terrestrial resources for food complemented, in some A definitive rupture with foraging subsistence strate- coastal areas, with marine organisms. Considerable isotopic gies appears to have occurred during the Late Neolithic– variability among Mesolithic individuals could be associ- Chalcolithic. The narrow isotopic distribution of individ- ated with versatile and diversified subsistence strategies, ual diets, in particular in the Mediterranean region, sug- adapted to the mosaic of environmental conditions in the gests an intensification of farming practices in an Iberian Peninsula. 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Routledge, New York, pp 53–73 Bogaard A, Heaton THE, Poulton P, Merbach I (2007) The impact of Acknowledgements This paper is a contribution to the Marie manuring on nitrogen isotope ratios in cereals: archaeological impli- Sklodowska Curie research project First ceramics of Atlantic Europe: cations for reconstruction of diet and crop management practices. J manufacture and function (CerAM -IF 653354-) funded by the Archaeol Sci 34:335–343. https://doi.org/10.1016/j.jas.2006.04.009 European Commission. The authors wish to thank Peter Smith and Bogaard A, Frase R, Heaton T, Wallace M, Vaiglova P, Charles M, Jones Krista McGrath for their review of the manuscript. In addition, we would G, Evershed R, Styring A, Andersen N, Arbogast RM, Bartosiewicz like to thank the two anonymous reviewers for their suggestions and L, Gardeisen A, Kanstrup M, Maier U, Marinova E, Ninov L, comments. Schäfer M, Stephan E (2013) Crop manuring and intensive land management by Europe’s first farmers. Proc Natl Acad Sci 110: Funding information Miriam Cubas is supported by Fellows Gipuzkoa 12589–12594. https://doi.org/10.1073/pnas1305918110 Research Program funded by the Diputación Foral de Gipuzkoa. Bonsall C, Cook GT, Pickard C, McSweeney K, Bartosiewicz L (2009) Dietary trends at the Mesolithic-Neolithic transition in North-West Europe. In: Crombé P, Van Strydonck M, Sergant J, Bioudin M, Bats Publisher’sNote Springer Nature remains neutral with regard to juris- M (eds) Chronology and evolution within the Mesolithic of North- dictional claims in published maps and institutional affiliations. West Europe. Cambridge Scholars Publishing, Newcastle upon Tyne, pp 539–562 Open Access This article is distributed under the terms of the Creative Carvalho AF, Petchey F (2013) Stable isotope evidence of Neolithic Commons Attribution 4.0 International License (http:// paleodiets in the coastal regions of southern Portugal. 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